Associations Among Breeding Birds and Gambel Oak in Southwestern Ponderosa Pine Forests

Abstract: Ponderosa pine (Pinus ponderosa) forests with Gambel oak (Quercus gambelii) are associated with higher bird abundance and diversity than are ponderosa pine forests lacking Gambel oak. Little is known, however, about specific structural characteristics of Gambel oak trees, clumps, and stands that may be important to birds in ponderosa pine-Gambel oak (hereafter pine-oak) forests. We examined associations among breeding birds and structural characteristics of Gambel oak at a scale similar in size to individual bird territories in pine-oak forests in northern Arizona and western New Mexico, USA. Avian species richness and occurrence of some bird species were associated with specific growth forms of Gambel oak. Estimated probability of Virginia's warblers (Vermivora virginiae), black-headed grosbeaks (Pheucticus melanocephalus), and red-faced warblers (Cardellina rubrifrons) occurring at points increased with increasing density of pole-sized Gambel oak 7–15 cm in diameter at breast height. We also found evidence that large Gambel oak trees (≥23 cm dbh) were associated with increased occurrence of yellow-rumped warblers (Dendroica coronata) at points. Some avian associations with oak were influenced by characteristics of ponderosa pines. For example, bird species richness was positively associated with the abundance of large Gambel oak when density of large pine trees ≥23 cm in diameter at breast height was low. Because large oak trees are rare and their numbers are thought to be declining, efforts should be made to retain and promote growth of additional oaks in this size class. Forest management practices that maintain forest openings, such as prescribed burning, could promote growth of pole-sized Gambel oak, which appears important to some bird species in pine-oak forests.     

The African rainforest: odd man out or megafaunal landscape? African and Amazonian forests compared

Africa has been called the ‘odd man out’ because the hectare‐scale tree diversity of African equatorial forests is lower than that of forests in other parts of the tropics. Low diversity has been attributed to the smaller area of the African forest and a history of drought, fire and contraction. Several facts shed doubt on this interpretation. The current area of the central African forest is roughly 2 million km². Even during periods of Pleistocene contraction, numerous moist refugia remained, including 6 posited for Gabon, a country the size of the U. S. state of Colorado. The gamma‐diversity of Gabon is high, implying higher alpha diversities. Finally, tree diversities on small islands in the Solomons and Fiji archipelagos are twice those prevalent in Gabonese forests, suggesting that historical contractions may not have been sufficient to reduce diversity to its current level. To place the African situation in perspective, we compared tree stands in Gabon and the Peruvian Amazon. Peruvian forests contained a mean of 618 trees ≥ 10 cm dbh per ha vs 377 for Gabon, or 64% more. Peruvian forests contained relatively more small trees (≥ 10, <20 cm dbh) and many fewer large trees (≥ 20 cm dbh) than Gabonese forests. These structural differences were consistent across 10 Gabonese and 10 Peruvian sites and transcended local gradients in climate and geology, suggesting that they are intrinsic to the two continents. Tree species diversity in Perú is concentrated in the small tree class (≥ 10, <20 cm dbh), whereas it is highest in the larger tree classes in Gabon. Alpha diversity is apparently lower relative to gamma diversity in Africa than it is in Amazonian Perú , implying higher beta diversity. The densities of small plants (<1 m tall) are similar in Gabonese and Peruvian forests; the observed structural differences develop later at the sapling and small tree stages. Explaining the low hectare‐scale diversity of African forests thus reduces to understanding why the density and diversity of small trees is so anomalously low.     

Forest ecosystems of an Arizona Pinus ponderosa landscape: multifactor classification and implications for ecological restoration

Aim We developed an ecosystem classification within a 110,000‐ha Arizona Pinus ponderosa P. & C. Lawson (ponderosa pine) landscape to support ecological restoration of these forests. Specific objectives included identifying key environmental variables constraining ecosystem distribution and comparing plant species composition, richness and tree growth among ecosystems. Location The Coconino National Forest and the Northern Arizona University Centennial Forest, in northern Arizona, USA. Methods We sampled geomorphology, soils and vegetation on 66 0.05‐ha plots in open stands containing trees of pre‐settlement (c. 1875) origin, and on 26 plots in dense post‐settlement stands. Using cluster analysis and ordination of vegetation and environment matrices, we classified plots into ecosystem types internally similar in environmental and vegetational characteristics. Results We identified 10 ecosystem types, ranging from dry, black cinders/Phacelia ecosystems to moist aspen/Lathyrus ecosystems. Texture, organic carbon and other soil properties reflecting the effects of parent materials structured ecosystem distribution across the landscape, and geomorphology was locally important. Plant species composition was ecosystem‐specific, with C₃Festuca arizonica Vasey (Arizona fescue), for instance, abundant in mesic basalt/Festuca ecosystems. Mean P. ponderosa diameter increments ranged from 2.3–4.3 mm year^?1 across ecosystems in stands of pre‐settlement origin, and the ecosystem classification was robust in dense post‐settlement stands. Main conclusions Several lines of evidence suggest that although species composition may have been altered since settlement, the same basic ecosystems occurred on this landscape in pre‐settlement forests, providing reference information for ecological restoration. Red cinders/Bahia ecosystems were rare historically and > 30% of their area has been burned by crown fires since 1950, indicating that priority could be given to restoring this ecosystem's remaining mapping units. Ecosystem classifications may be useful as data layers in gap analyses to identify restoration and conservation priorities. Ecosystem turnover occurs at broad extents on this landscape, and restoration must accordingly operate across large areas to encompass ecosystem diversity. By incorporating factors driving ecosystem composition, this ecosystem classification represents a framework for estimating spatial variation in ecological properties, such as species diversity, relevant to ecological restoration.     

Characteristics of snags and forest structure in southern mistbelt forests of the Amatole region,South Africa

Standing dead trees (or snags) are an important component of forest ecosystems, especially for tree cavity‐nesting vertebrate species, but their prevalence in South African forests remains under studied. Consequently, we investigated forest structure, and the presence and abundance of snags in six southern mistbelt forests in the Eastern Cape, South Africa. These forests have had varying levels of timber extraction over the past 150 years or more. We found snags were relatively rare in all six forests (<4.3% of trees sampled). Mean diameter at breast height (dbh) of snags ranged from 52 to 82 cm across the forests, with smaller snags in Kologha Forest and larger snags in Tyume Forest. A bimodal distribution of snag successional stages was found, with frequencies peaking at early and late stages, and few in the intermediate stages. Tree species diversity in the forests was relatively low (twelve–nineteen species across forests; only 28 species in total). There was no significant difference in dbh of trees between forests, with most occurring in the 20–29‐cm dbh size class. Future studies are required to identify trees that most likely support suitable cavities for tree cavity‐nesting bird species, and to determine cavity‐nester assemblage requirements in southern African forests.     

Effects of Restoration Thinning on Presettlement Pinus ponderosa in Northern Arizona

In the 100 years following the arrival of Euro-American settlers in northern Arizona, Pinus ponderosa (ponderosa pine) forests changed from open, low-density stands to closed, high-density stands. The increase in tree density has been detrimental to the vigor of old-growth trees that established before settlement (presettlement trees). In this study, we examined whether the vigor of presettlement trees could be improved by restoring the original stand structure by thinning the ponderosa pines that established after settlement (postsettlement trees). The restoration treatment caused the following changes in the presettlement trees and their environment in the first year following thinning: an increase in volumetric soil water content between May and August, an increase in predawn xylem water potential in July and August, a decrease in midday xylem water potential in June and August, an increase in net photosynthetic rate in August, an increase in foliar nitrogen concentration in July and August, and an increase in bud and needle size. The results show that the thinning restoration treatment improved the condition of presettlement ponderosa pines by increasing canopy growth and the uptake of water, nitrogen, and carbon.     

Monitoring Landscape‐Scale Ponderosa Pine Restoration Treatment Implementation and Effectiveness

We evaluated landscape‐scale forest restoration treatment implementation and effectiveness in meeting objectives in a ponderosa pine forest at Mt. Trumbull, Arizona, U.S.A. The goal of the project was to alter forest structure by thinning and burning to more closely resemble forest conditions prior to Euro‐American settlement in 1870. We measured 117 permanent plots before (1996/1997) and after (2003) treatments. The plots were evenly distributed across the landscape (approximately 1,200 ha), about half of which was an untreated control. We evaluated treatment implementation and effectiveness based on 1870 structure and/or goals outlined by managers. The success of treatment implementation varied: about 94% of the area originally planned for restoration was treated in some manner by 2003, but only 70% received the full planned treatment (thin and burn). Although density of ponderosa pines >2.5 cm was reduced significantly by 66% from approximately 429 pines/ha to approximately 146 pines/ha in the treated area, the targeted residual density was exceeded by 111–256% (all plots) or 10–85% (thinned and burned plots). Thirteen percent of the pre‐settlement pines died in the treated area by 2003, but 9% percent also died in the control, indicating that pre‐settlement pines in untreated areas were nearly as vulnerable as those exposed to restoration treatments. Large snags increased 45%, and 65% of logs >50 cm were retained, achieving implementation goals. Although restoration treatments were not implemented totally to specifications, they were effective in attaining the overall project goal of restoring more open forest structure while preserving more than 75% of the pre‐settlement pines. Canopy fuel loads were substantially reduced, allowing for the reintroduction of surface fires.     

Liana cover in the canopies of rainforest trees is not predicted by local ground‐based measures

Lianas (woody climbers) are structural parasites of trees that compete with them for light and below‐ground resources. Most studies of liana–tree interactions are based on ground‐level observations of liana stem density and size, with these assessments generally assumed to reflect the amount of liana canopy cover and overall burden to the tree. We tested this assumption in a 1‐ha plot of lowland rainforest in tropical Australia. We recorded 1072 liana stems (≥1 cm diameter at breast height {dbh}) ha^?1 across all trees (≥10 cm dbh) on the plot and selected 58 trees for detailed study. We estimated liana canopy cover on selected trees that hosted 0–15 liana individuals, using a 47‐m‐tall canopy crane. Notably, we found no significant correlations between liana canopy cover and three commonly used ground‐based measurements of liana abundance as follows: liana stem counts per tree, liana above‐ground biomass per tree and liana basal area per tree. We also explored the role of tree size and liana infestation and found that larger trees (≥20 cm dbh) were more likely to support lianas and to host more liana stems than smaller trees (≤20 cm dbh). This pattern of liana stem density, however, did not correlate with greater liana coverage in the canopy. Tree family was also found to have a significant effect on likelihood of hosting lianas, with trees in some families 3–4 times more likely to host a liana than other families. We suggest that local ground‐based measures of liana–tree infestation may not accurately reflect liana canopy cover for individual trees because they were frequently observed spreading through neighbouring trees at our site. We believe that future liana research will benefit from new technologies such as high‐quality aerial photography taken from drones when the aim is to detect the relative burden of lianas on individual trees.     

The Relationship between Local Abundance and Distribution of Rain Forest Trees across Environmental Gradients in India

We tested whether local abundance of rain forest trees in the medium elevation wet forests of the southern Western Ghats (WG) was related to environmental tolerance, life form, and geographical range. We selected trees in medium elevation wet forests (750–1700 m asl) of the southern WG, using two data bases: a small plot (30 × 30 m) data base of 288 species of trees (≥ 3 cm dbh) in 33 plots totaling 2.97 ha, and a data base of 135 species of tree (≥ 10 cm dbh) in larger plots of 1 ha each, totaling 4.84 ha. The species density per hectare and number of records in the plot network was used in a factor analysis to give a measure of the local abundance of each species. The altitude and seasonality ranges of these species in the WG was assessed from independent data bases and used to generate an environmental tolerance score. Results indicated that as a species became locally more abundant, it occurred across a wider range of environmental gradients, but regional distribution was not related to geographical distribution. Understory species tended to be rarer with smaller range sizes and lower environmental tolerances than overstory species. Climate change is predicted to have drastic effects on restricted range species with limited environmental tolerances.     

Reference conditions and historical changes in an unharvested ponderosa pine stand on sedimentary soil

Much of the previous research on spatial reference conditions in dry frequent fire pine forests have come from stand‐level patterns under regionally average ecosystem conditions (e.g. soil type and precipitation). We evaluated the 1883 reference conditions of an uncut ponderosa pine stand representing a far end of the range of variability in terms of regionally unusual environmental conditions. Using a forest reconstruction model, univariate and bivariate Ripley's K functions, and regression analysis, we determined 1883 structural and spatial reference conditions, and compared those to the contemporary (2010) stand. Historical stand density was 77 trees/ha with a basal area of 8.0 m²/ha. Reference spatial patterns were significantly aggregated from 1 to 2 m and randomly distributed at distances greater than 2 m. Nearly 40% of the reconstructed trees were individuals, the average patch size was 2.9 trees, and the largest patch had 7 members. The contemporary stand had considerably greater densities and basal area than historical conditions and showed aggregation at all distances. Bivariate spatial analysis indicated attraction of post‐settlement recruitment to live pre‐settlement trees from 1 to 6 m and no association at distances greater than 6 m. We speculate that the historically random tree pattern is the product of a variety of factors including soil parent material, climate, and more homogeneous resource partitioning.     

Tree population studies in low-diversity forests,Guyana. I. Floristic composition and stand structure

Studies were undertaken on the floristic composition and stand structure of four 1 hectare plots in the lowland forests of Kurupukari, Guyana. A total of 3897 trees, covering 153 species and 31 plant families were recorded at greater than 5 cm diameter at breast height (dbh). The number of species per hectare ranged from 61 to 84 (>5.0 cm dbh) and 50–71 (>10.0 cm dbh). The total number of trees per hectare varied two-fold between study plots, with 45–50% of the trees within the 5–10 cm size-class. Mean total basal area varied from 32.39–34.63 m² per 100 m². The four most dominant plant families represented 43.8% of the total number of trees, while representing only 11.2% of the species. No one plant family dominated in more than one of the four study plots, and all four plots held at least one plant family with more than 20% of the total number of trees. Although 14 tree species were common to all four plots, only 26%–35% of the species were represented by a single tree. Between three and seven species represented 50% of the trees within all size-classes, with species dominance occurring within the highest density plot.These tropical forest types of central Guyana may represent some of the lowest diversity forests in the neotropics, whereby the total number of tree species is relatively limited, typically with six dominant canopy species, but the relative abundance of these species is highly variable between the forest types. Mechanisms influencing the competitive interactions associated with species dominance are discussed in relation to the importance of mycorrhizae and the persistence of species dominance.     

Diameter Growth of Juvenile Trees after Gap Formation in a Bolivian Rain Forest: Responses are Strongly Species‐specific and Size‐dependent

We evaluated growth responses to gap formation for juvenile individuals of three canopy rain forest species: Peltogyne cf. heterophylla, Clarisia racemosa and Cedrelinga catenaeformis. Gaps were formed during selective logging operations 7 yr before sampling in a Bolivian rain forest. We collected wood samples for tree‐ring analyses at different distances to the stump (<10, 10–40 and >40 m) and from trees with different diameters (5–30 cm diameter at breast height [dbh]). Tree‐rings width was measured in at least two radii and converted to average diameter growth. Changes in 7‐yr median diameter growth before and after selective logging were analyzed. Diameter growth rates significantly increased by 0.7–0.8 mm/yr after gap formation for P. heterophylla and C. catenaeformis, but not for C. racemosa. We applied a multiple regression analysis to explain variation in growth responses of P. heterophylla and C. catenaeformis by distance to logging gap and tree size. For P. heterophylla we found that growth increase occurring close to logging gaps was strongest for large juvenile trees (20–25 cm dbh) and almost absent in small juveniles. For C. catenaeformis, variation in growth responses was not related to tree size or distance to gaps. Our results show that growth responses to gap formation strongly differ across species and tree sizes. This finding calls for caution in the interpretation of growth releases in tree‐ring series, as gap formation does not necessarily invoke growth responses and if such growth responses occur, their strength is species‐ and size specific.     

Fire, fuels and restoration of ponderosa pine–Douglas fir forests in the Rocky Mountains, USA

Aim Forest restoration in ponderosa pine and mixed ponderosa pine–Douglas fir forests in the US Rocky Mountains has been highly influenced by a historical model of frequent, low‐severity surface fires developed for the ponderosa pine forests of the Southwestern USA. A restoration model, based on this low‐severity fire model, focuses on thinning and prescribed burning to restore historical forest structure. However, in the US Rocky Mountains, research on fire history and forest structure, and early historical reports, suggest the low‐severity model may only apply in limited geographical areas. The aim of this article is to elaborate a new variable‐severity fire model and evaluate the applicability of this model, along with the low‐severity model, for the ponderosa pine–Douglas fir forests of the Rocky Mountains. Location Rocky Mountains, USA. Methods The geographical applicability of the two fire models is evaluated using historical records, fire histories and forest age‐structure analyses. Results Historical sources and tree‐ring reconstructions document that, near or before ad 1900, the low‐severity model may apply in dry, low‐elevation settings, but that fires naturally varied in severity in most of these forests. Low‐severity fires were common, but high‐severity fires also burned thousands of hectares. Tree regeneration increased after these high‐severity fires, and often attained densities much greater than those reconstructed for Southwestern ponderosa pine forests. Main conclusions Exclusion of fire has not clearly and uniformly increased fuels or shifted the fire type from low‐ to high‐severity fires. However, logging and livestock grazing have increased tree densities and risk of high‐severity fires in some areas. Restoration is likely to be most effective which seeks to (1) restore variability of fire, (2) reverse changes brought about by livestock grazing and logging, and (3) modify these land uses so that degradation is not repeated.     

Population structure and dynamics oi Juniperus excelsa in Balouchistan,Pakistan

Abstract. 60 monospecific stands of Juniperus excelsa were sampled at four locations in Balouchistan. Density, basal area and height of individuals were recorded. Soils were analysed for selected physical and chemical characteristics and the degree of disturbance due to logging and burning was noted. The density of juniper trees (> 6 cm dbh) ranged from 56 to 332 stems / ha (average 174 stems / ha). Higher densities were recorded for relatively undisturbed stands and on west facing slopes. Density of seedlings and saplings (< 6 cm dbh) was strongly correlated with tree density and tree basal area. Among the edaphic variables CaC0₃ was correlated with juniper density and basal area. Diameter distributions within stands were mostly skewed and unimodal with gaps appearing in large size classes. The male to female ratio was close to 1. Cross-sections of 16 trees were used to determine age and growth rate. Number of rings in trees with 20 to 30 cm dbh ranged from 95 to 221 (x = 160 ± 38). Diameter and age were not related. Mean annual diameter increment ranged from 6 to 16 yr / cm x = 10 ± 3 yr / cm). It is concluded that size class gaps and low seedling / sapling densities are the consequence of anthropogenic disturbance.     

Water and Nutrient Outflow Following the Ecological Restoration of a Ponderosa Pine-Bunchgrass Ecosystem

In the late 1800s, fire suppression, livestock grazing, and a wet and warm climate led to an irruption of pine regeneration in Pinus ponderosa Laws. (ponderosa pine) forests of the southwestern United States. Pines invaded bunchgrass openings, causing stand structure changes that increased the number of stand-replacing fires. Ecological restoration, via thinning and prescribed burning, is being used to decrease the risk of stand-replacing fires and ameliorate other effects of pine invasion. The effects of aboveground restoration on belowground processes are poorly understood. We used a hydrologic model and soil water nutrient concentrations, measured monthly below the rooting zone, to estimate restoration effects on nutrient losses by leaching from a mature ponderosa pine forest near Flagstaff, Arizona. Replicated restoration treatments included thinning to pre-1880 stand densities (partial restoration), thinning plus forest floor fuel reduction followed by a prescribed burn (complete restoration), and an untreated control. Water outflow occurred only between January and May and was lowest from the control (47 and 28 mm in 1995 and 1996) and highest from the partial restoration treatment (67 and 59 mm in 1995 and 1996). The concentrations (typically <0.10 mg/ L) and estimated annual losses (<0.02 kg/ha) of NH₄⁺-N, PO₄^{3 ?} -P, and organic P were similar among treatments. Nitrate and organic N concentrations were as high as 0.80 mg N/L; however, these concentrations and estimated annual losses (<0.13 kg N/ha) were similar among treatments. Our results suggest that restoration will not enhance nutrient loss by leaching or alter stream chemistry in ponderosa pine forests.     

Structure and Composition of Historical Longleaf Pine Ecosystems in Mississippi,USA

Longleaf pine (Pinus palustris) historically was a widespread ecosystem composed of a simple tree canopy and grasslands ground layer. After widespread loss of this ecosystem due to logging and fire exclusion, little quantitative information exists about historical structure for restoration goals. We identified composition in De Soto National Forest and Pearl River County, Mississippi, USA, and density, basal area, and percent stocking in Pearl River County using General Land Office surveys and US Forest Service Forest Inventory and Analysis surveys. Historical longleaf ecosystems were about 85% pine, with lesser amounts of broadleaf evergreen and oak species. Densities were about 175 to 180 trees/ha, mean tree diameters were 45 cm, and stocking was around 60% to 65%, which suggested longleaf pines were closed woodlands. Current forests are 38% to 57% pine, primarily loblolly, while longleaf pine is 2% to 8% of composition. Indeed, current longleaf pine composition across the Coastal Plain averages 3% and does not reach 10% at smaller landscape scales. Fire-sensitive broadleaf species of water oak, sweetgum, yellow-poplar, and red maple increased from about 0.5% composition to 2% to 10% of composition. Forests became twice as dense, at about 280 trees/ha to 330 trees/ha, with mean tree diameters of 22 cm. These results characterize conversion from open old growth longleaf forests, resulting in part from human maintenance, to successional forests due to human disruption of the historical ecosystem. It is important to remember structure and composition of historical forests for restoration and recognize wholesale changes so that successional forests do not become the new social and cultural baseline.     

Modeling Ecological Restoration Effects on Ponderosa Pine Forest Structure

FIRESUM, an ecological process model incorporating surface fire disturbance, was modified for use in southwestern ponderosa pine ecosystems. The model was used to determine changes in forest structure over time and then applied to simulate changes in aboveground biomass and nitrogen storage since exclusion of the natural frequent fire regime in an unharvested Arizona forest. Dendroecological reconstruction of forest structure in 1876, prior to Euro‐American settlement, was used to initialize the model; projections were validated with forest measurements in 1992. Biomass allocations shifted from herbaceous plants to trees, and nitrogen was increasingly retained in living and dead tree biomass over the 116‐year period (1876–1992). Forest conditions in 1992 were substantially degraded compared to reference presettlement conditions: old‐growth trees were dying at accelerated rates, herbaceous production was reduced nearly 90%, and the entire stand was highly susceptible to high‐intensity wildfire. Following an experiment initiated in 1993 to test ecological restoration treatments, future changes were modeled for the next century. Future forest structure remained within the natural presettlement range of variability under the full restoration treatment, in which forest biomass structure was thinned to emulate presettlement conditions and repeated low‐intensity fire was reintroduced. Simulation of the control treatment indicated continuation of exceptionally high tree density, probably culminating in stand‐replacing ecosystem change through high‐intensity wildfire or tree mortality from pathogens. Intermediate results were observed in the partial restoration treatment (tree thinning only); the open forest structure and high herbaceous productivity found immediately after treatment were gradually degraded as dense tree cover reestablished in the absence of fire. Modeling results support comprehensive restorative management as a long‐term approach to conservation of key indigenous ecosystem characteristics.     

Importance of resin ducts in reducing ponderosa pine mortality from bark beetle attack

The relative importance of growth and defense to tree mortality during drought and bark beetle attacks is poorly understood. We addressed this issue by comparing growth and defense characteristics between 25 pairs of ponderosa pine (Pinus ponderosa) trees that survived and trees that died from drought-associated bark beetle attacks in forests of northern Arizona, USA. The three major findings of our research were: (1) xylem resin ducts in live trees were >10% larger (diameter), >25% denser (no. of resin ducts mm⁻²), and composed >50% more area per unit ring growth than dead trees; (2) measures of defense, such as resin duct production (no. of resin ducts year⁻¹) and the proportion of xylem ring area to resin ducts, not growth, were the best model parameters of ponderosa pine mortality; and (3) most correlations between annual variation in growth and resin duct characteristics were positive suggesting that conditions conducive to growth also increase resin duct production. Our results suggest that trees that survive drought and subsequent bark beetle attacks invest more carbon in resin defense than trees that die, and that carbon allocation to resin ducts is a more important determinant of tree mortality than allocation to radial growth.